Kent Academic Repository Full text document (pdf) Citation for published version Zanolli, Clément and Biglari, Fereidoun and Mashkour, Marjan and Abdi, Kamyar and Monchot, Hervé and Debue, Karyne and Mazurier, Arnaud and Bayle, Priscilla and Le Luyer, Mona and Rougier, Hélène and Trinkaus, Erik and Macchiarelli, Roberto (2019) A Neanderthal from the Central Western Zagros, Iran. Structural reassessment of the Wezmeh 1 maxillary premolar. DOI Link to record in KAR https://kar.kent.ac.uk/75685/ Document Version Author's Accepted Manuscript Copyright & reuse Content in the Kent Academic Repository is made available for research purposes. Unless otherwise stated all content is protected by copyright and in the absence of an open licence (eg Creative Commons), permissions for further reuse of content should be sought from the publisher, author or other copyright holder. Versions of research The version in the Kent Academic Repository may differ from the final published version. Users are advised to check http://kar.kent.ac.uk for the status of the paper. Users should always cite the published version of record. Enquiries For any further enquiries regarding the licence status of this document, please contact: [email protected] If you believe this document infringes copyright then please contact the KAR admin team with the take-down information provided at http://kar.kent.ac.uk/contact.html A Neanderthal from the Central Western Zagros, Iran. Structural reassessment of the Wezmeh 1 maxillary premolar Clément Zanolli a, *, Fereidoun Biglari b, *, Marjan Mashkour c, d, Kamyar Abdi e, Hervé Monchot f, Karyne Debue c, Arnaud Mazurier g, Priscilla Bayle a, Mona Le Luyer a, h, Hélène Rougier i, Erik Trinkaus j, Roberto Macchiarelli k,l a Laboratoire PACEA, UMR 5199 CNRS, Université de Bordeaux, Ministère de la Culture, Bordeaux, France b Paleolithic Department, National Museum of Iran, Tehran, Iran c Laboratoire AASPH, UMR 7209 CNRS, Muséum national d'Histoire naturelle, Paris, France d Archaeozoology Section, Bioarchaeology Laboratory, Central Laboratory, University of Tehran, Tehran, Iran e University of Shiraz, Shiraz, Iran f UMR 8167 Orient et Méditerranée, Paris Sorbonne Universités, Paris, France g Institut de Chimie des Milieux et Matériaux, UMR 7285 CNRS, Université de Poitiers, Poitiers, France h Skeletal Biology Research Centre, School of Anthropology and Conservation, University of Kent, Canterbury, UK i Department of Anthropology, California State University, Northridge, CA 91330, USA j Department of Anthropology, Washington University, Saint Louis, MO 63130, USA k Laboratoire HNHP, UMR 7194 CNRS, Muséum national d'Histoire naturelle, Musée de l'Homme, Paris, France l Unité de Formation Géosciences, Université de Poitiers, Poitiers, France * Corresponding authors. E-mail addresses: [email protected] (C. Zanolli), [email protected] (F. Biglari). Acknowledgments Research funded by the French CNRS. We thank the National Museum of Iran for research support, contribution and generous collaboration; the UMR 7209 for funding the surface scans at the SURFAçUS Platform of the MNHN Paris and the 3D print of Wezmeh 1; the PLATeforme INstrumentale d’Analyse of the IC2MP laboratory of the University of Poitiers for access to the µCT device; and the LabEx "Sciences archéologiques" of the University of Bordeaux for the analysis of comparative materials. We also thank the Editor, the Associate Editor, and the two anonymous reviewers for their helpful comments on the manuscript. 1 A Neanderthal from the Central Western Zagros, Iran. Structural reassessment of 2 the Wezmeh 1 maxillary premolar 3 4 ABSTRACT 5 6 Wezmeh Cave, in the Kermanshah region of Central Western Zagros, Iran, produced a Late 7 Pleistocene faunal assemblage rich in carnivorans along with a human right maxillary premolar, 8 Wezmeh 1, an unerupted tooth from an 8 ± 2 year-old individual. Uranium-series analyses of the 9 fauna by alpha spectrometry provided age estimates between 70 and 11 ka. Crown dimensions 10 place the tooth specimen at the upper limits of Late Pleistocene human ranges of variation. Wezmeh 11 1 metameric position (most likely a P3) remains uncertain and only its surficial morphology has 12 been described so far. Accordingly, we used micro-focus X-ray tomography (12.5 µm isotropic 13 voxel size) to reassess the metameric position and taxonomic attribution of this specimen. We 14 investigated its endostructural features and quantified crown tissue proportions. Topographic maps 15 of enamel thickness (ET) distribution were also generated, and semilandmark-based geometric 16 morphometric analyses of the enamel-dentine junction (EDJ) were performed. We compared 17 Wezmeh 1 with unworn/slightly-moderately worn P3 and P4 of European Neanderthals, Middle 18 Paleolithic modern humans from Qafzeh, an Upper Paleolithic premolar, and Holocene humans. 19 The results confirm that Wezmeh 1 represents a P3. Based on its internal conformation and 20 especially EDJ shape, Wezmeh 1 aligns closely with Neanderthals and is distinct from the fossil 21 and extant modern human pattern of our comparative samples. Wezmeh 1 is thus the first direct 22 evidence of Neanderthal presence on the western margin of the Iranian Plateau. 23 1 24 Keywords: Zagros Mountains; Wezmeh Cave; Human premolar tooth; Microfocus X-ray 25 tomography; Neanderthals 26 2 27 1. Introduction 28 Since Coon’s pioneering excavation in the late 1940s at the Middle Paleolithic site of Bisitun 29 Cave (Coon, 1951), the Zagros Mountains of Iran and Iraq have produced a number of Late 30 Pleistocene human remains associated with Middle and Upper Paleolithic technocultural contexts 31 that represent the main bioanthropological source of information about the populations inhabiting 32 the region during the oxygen isotope stages (OIS) 3–2 (for a recent chronometric assessment of 33 some Middle and Upper Paleolithic sites of the region, see Becerra-Valdivia et al., 2017). Among 34 the excavated sites yielding human remains, with its assemblage of at least ten Neanderthal 35 individuals, the Shanidar Cave, Iraqi Kurdistan, has produced one of the largest sample of fossil 36 human remains from Southwestern Asia thus far (Solecki, 1963; Trinkaus, 1983; Cowgill et al., 37 2007; Pomeroy et al., 2017). 38 A complex of Paleolithic sites in the Kermanshah region of Central Western Zagros, Iran, 39 produced the second fossil human sample (Fig. 1). Among these sites, the Middle Paleolithic 40 sequence of Bisitun Cave, located east-northeast of Kermanshah, was the first that yielded a human 41 fossil (Coon, 1951), a right radius preserved from just distal of the radial tuberosity to the distal 42 diaphysis. Although its dimensions overlap the upper end of the earlier Upper Paleolithic range of 43 variation, comparisons with Neanderthal and Middle Paleolithic early modern human right radii 44 align it predominantly with Neanderthals, but separate from Middle Paleolithic early modern 45 humans of Southwestern Asia (Trinkaus and Biglari, 2006). 46 In the course of a reanalysis at the Field Museum of Natural History of the Upper Paleolithic 47 collections from the Warwasi rockshelter, a site in the northern outskirt of Kermanshah, an isolated 48M 2 from a fossil modern human was recently identified in the faunal assemblage from the 49 Baradostian levels (Tsanova, 2013). 3 50 Another Paleolithic site in Western Zagros that provided human remains is Gar Arjeneh, near 51 Khoramabad, in Luristan. First excavated by Hole and Flannery (1967; for a review see Bazgir et 52 al., 2014), its Baradostian levels have yielded a human premolar and some bone fragments 53 (Trinkaus, 2018). 54 In the Southern Zagros, the Upper Paleolithic and Epipaleolithic layers of Eshkaft-e Gavi, near 55 Shiraz, produced an assemblage of fragmentary dental, cranial and postcranial remains sampling a 56 minimum of four modern human individuals bearing evidence of stone-tool cutmarks and burning 57 (Rosenberg, 1985; Scott and Marean, 2009). 58 Finally, the catalog of human remains from Paleolithic sites of Central Western Zagros 59 Mountains includes Wezmeh 1, an isolated maxillary premolar from Wezmeh Cave, southwest of 60 Kermanshah. Its buccolingual crown diameter places it at the upper limits of Late Pleistocene 61 human maxillary third (P3) and fourth premolar (P4) dimensions and above a terminal Pleistocene 62 Natufian sample (Trinkaus et al., 2008). 63 Given uncertainties concerning the original chronostratigraphic context of Wezmeh 1 (Abdi et 64 al., 2002; Trinkaus et al., 2008; Mashkour et al., 2009) and the difficulties of assessing its 65 taxonomic status based on its external features (Trinkaus et al., 2008), we used micro-focus X-ray 66 tomography (µCT) to reassess the metameric position and taxonomic attribution of this specimen. 67 Using techniques of virtual imaging coupled with geometric morphometrics, we evaluated its tissue 68 proportions, enamel thickness topographic distribution, enamel-dentine junction morphology, and 69 pulp cavity shape. We then compared these data to samples of European Neanderthals, Near 70 Eastern Middle Paleolithic modern humans, an Upper Paleolithic human, and Holocene humans. 71 72 1.1. The Wezmeh Cave 4 73 Wezmeh Cave is located 10 km southeast of the town of Islamabad-e Gharb (34°03'17.04" N, 74 46°38'41.03" E), at an elevation of ca. 1430 m asl (Fig. 1). The cave, discovered in 1999 during 75 archeological survey as a part of the Islamabad Project (director: K.A.), is in a small valley in the 76 western foothills of the Qazivand Mountain (Fig. 2a). Its entrance (Fig. 2b) faces north and is 2 m 77 wide and 1.2 m high. The cave is about 27 m long and has about 45 m2 of floor area (Abdi et al., 78 2002; see Fig. 2c). In 2001, following disturbance by clandestine digging, a 3 × 3 m trench was 79 excavated to the bedrock on the terrace immediately outside of the cave entrance, and six small test 80 pits at 2 m intervals were dug to bedrock within the cave.